Habitat light, colour variation, and ultraviolet reflectance in the Grand Cayman anole, Anolis conspersus
01 Jul 2001-Biological Journal of The Linnean Society (No longer published by Elsevier)-Vol. 73, Iss: 3, pp 299-320
TL;DR: Evidence is presented to show how geological, ecological, and physiological factors could have interacted to select for a short wavelength-reflective dewlap from a long wavelength- reflective precursor following the colonization of Grand Cayman from Jamaica by A. grahami between 2 and 3 Mya.
Abstract: Data from a diversity of sources are consistent with the hypothesis that the Grand Cayman anole, Anolis conspersus, is descended directly from Anolis grahami of Jamaica. Although the two species have remained morphologically similar, coloration in A. conspersus has changed considerably from that of its ancestor. The most dramatic difference is seen in dewlap colour, where A. conspersus has evolved a blue and highly UV-reflective dewlap from the ancestral orange-and-yellow colour state. In addition, variation in normal (non-metachrosis) dorsum coloration in A. grahami populations is limited to shades of green (olive, emerald, teal), whereas in A. conspersus dorsum coloration varies from green to blue and to brown. This increased colour variation occurs despite Grand Cayman being a small, relatively featureless island only 35 km in length. Results of this study suggest that ambient light differences associated with precipitation-related vegetation structure may have played an important role in the evolution of A. conspersus body colour variation. Evidence is presented to show how geological, ecological, and physiological factors could have interacted to select for a short wavelength-reflective dewlap from a long wavelength-reflective precursor following the colonization of Grand Cayman from Jamaica by A. grahami between 2 and 3 Mya.
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TL;DR: Specific, testable functional hypotheses are offered for the most common pigmentary and structural components of vertebrate colour patches and how multiple trait evolution theory can be applied to the components of single colour patches.
Abstract: Colour patches are complex traits, the components of which may evolve independently through a variety of mechanisms. Although usually treated as simple, two-dimensional characters and classified as either structural or pigmentary, in reality colour patches are complicated, three-dimensional structures that often contain multiple pigment types and structural features. The basic dermal chromatophore unit of fishes, reptiles and amphibians consists of three contiguous cell layers. Xanthophores and erythrophores in the outermost layer contain carotenoid and pteridine pigments that absorb short-wave light; iridophores in the middle layer contain crystalline platelets that reflect light back through the xanthophores; and melanophores in the basal layer contain melanins that absorb light across the spectrum. Changes in any one component of a chromatophore unit can drastically alter the reflectance spectrum produced, and for any given adaptive outcome (e.g. an increase in visibility), there may be multiple biochemical or cellular routes that evolution could take, allowing for divergent responses by different populations or species to similar selection regimes. All of the mechanisms of signal evolution that previously have been applied to single ornaments (including whole colour patches) could potentially be applied to the individual components of colour patches. To reach a complete understanding of colour patch evolution, however, it may be necessary to take an explicitly multi-trait approach. Here, we review multiple trait evolution theory and the basic mechanisms of colour production in fishes, reptiles and amphibians, and use a combination of computer simulations and empirical examples to show how multiple trait evolution theory can be applied to the components of single colour patches. This integrative perspective on animal colouration opens up a host of new questions and hypotheses. We offer specific, testable functional hypotheses for the most common pigmentary (carotenoid, pteridine and melanin) and structural components of vertebrate colour patches.
245 citations
TL;DR: It is found that populations from mesic and xeric conditions occupy two distinct habitats with respect to light intensity and spectral quality and that dewlap design has diverged between populations in a way that increases signal detectability in each habitat.
Abstract: We tested the prediction of the sensory drive hypothesis using four allopatric populations of the lizard Anolis cristatellus from two distinct environments (i.e., mesic and xeric conditions). For each population, we measured habitat light characteristics and quantified signal design by measuring the spectral and total reflectance and transmittance of the dewlap. We used these data to calculate dewlap detectability using an empirically based model of signal detection probability. We found that populations from mesic and xeric conditions occupy two distinct habitats with respect to light intensity and spectral quality and that dewlap design has diverged between populations in a way that increases signal detectability in each habitat. The major difference in dewlap design was in total reflectance and transmittance, making dewlaps from xeric habitats darker and dewlaps from mesic habitats brighter. Furthermore, dewlap detection decreased significantly when a dewlap from a xeric habitat is detected u...
230 citations
Cites background from "Habitat light, colour variation, an..."
...Second, there is evidence that differences in habitat light conditions have selected for different color patterns in different microhabitats (i.e., selection through the process of sensory drive; Fleishman et al. 1993; Fleishman 2000; Macedonia 2001; Leal and Fleishman 2002)....
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...vol. 163, no. 1 the american naturalist january 2004 Differences in Visual Signal Design and Detectability between Allopatric Populations of Anolis Lizards Manuel Leal* and Leo J. Fleishman† Department of Biology, Union College, Schenectady, New York 12308...
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TL;DR: It is found that the chromatic and brightness contrasts of golden patches used during courtship are greater against the cleared court than against adjacent litter, and that cleared courts provide a less variable background for these color patches, resulting in displays that consistently contrast the visual background.
Abstract: Effective visual communication requires signals that are easy to detect, transmit, receive, and discriminate. Animals can increase the probability that their visual signals would be detected by evolving signals that contrast with their visual background. Animals can further enhance this contrast by behaviorally modifying the existing visual background. Male golden-collared manakins (Manacus vitellinus) clear leaf litter from the ground to form courts, which are used as display arenas. Using reflectance measures of the signal (male plumage) and the visual background (cleared court and adjacent litter), the irradiance measures of ambient light during display, and published measures of photoreceptor sensitivity of a Passerine, we test the hypothesis that court-clearing augments the contrast between male plumage and the visual background. We find that the chromatic and brightness contrasts of golden patches used during courtship are greater against the cleared court than against adjacent litter. In addition, we find that cleared courts provide a less variable background for these color patches, resulting in displays that consistently contrast the visual background. These results suggest that behavioral modification of the visual background may act to increase the conspicuousness of colorful male plumage during display, providing an explanation for why golden-collared manakins, and possibly other species, build or clear display courts. Key words: chromatic contrasts, court-clearing, Manacus vitellinus, manakins, signaling, visual signals. [Behav Ecol 15:1003–1010 (2004)]
146 citations
TL;DR: D dwarf chameleons showed different behavioural responses, including colour change, towards multiple predators that detect and capture prey in different ways, and whether these antipredator responses varied geographically was consistent among populations.
Abstract: Potential prey are often exposed to multiple predators that vary in their foraging tactics and ability to detect prey. For animals that rely on crypsis to avoid predators, one solution is to alter their behaviour or appearance to maximize crypsis in ways that are specific to different types of predator. We tested whether dwarf chameleons (Bradypodion transvaalense) showed different behavioural responses, including colour change, towards multiple predators (bird and snake models) that detect and capture prey in different ways, and whether these antipredator responses varied geographically. Chameleons consistently used the same body postures (lateral compression and flipping to the opposite side of the branch) and displayed similar chromatic (colour) contrast against the natural background in response to both predator types. However, they became significantly more achromatically contrasting (brighter) in the presence of the snake compared to the bird. This relative difference in achromatic contrast towards the two types of predator was consistent among populations. There were also significant differences in both absolute achromatic and chromatic contrast among populations despite very similar light environment, background coloration and habitat structure. Our results highlight facultative crypsis as one type of flexible antipredator tactic and emphasize the importance of visual ecology in understanding prey–predator interactions. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88, 437–446.
143 citations
Cites methods from "Habitat light, colour variation, an..."
...The contrast measures we used are simple measures of contrast in radiance (Endler, 1990; Macedonia, 2001; Heindl & Winkler, 2003)....
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...The contrast measures we used are simple measures of contrast in radiance (Endler, 1990; Macedonia, 2001; Heindl & Winkler, 2003)....
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...and background, respectively, integrated over 320‐ 700 nm. To calculate chromatic contrast, all radiance spectra of the chameleons and backgrounds were standardized for brightness (area under the curve 320‐ 700 nm = 1). Chromatic contrast was then calculated as the Euclidean distance ( D s ) between the radiance spectrum of the chameleon vs. background (Endler, 1990; Macedonia, 2001; Heindl & Winkler, 2003), using:...
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...Chromatic contrast was then calculated as the Euclidean distance ( D s ) between the radiance spectrum of the chameleon vs. background (Endler, 1990; Macedonia, 2001; Heindl & Winkler, 2003), using: where Q t ( λ ) and Q b ( λ ) is the radiance at a given wavelength and summation is over each 5 nm…...
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TL;DR: The results confirmed that there are real differences in crypsis conspicuousness both between populations and between sexes; that exposed body regions were significantly more cryptic than hidden ones, particularly in females; and that females, but not males, are more cryptic against their own local background than against the background of other populations.
Abstract: Many animal species display striking color differences with respect to geographic location, sex, and body region. Traditional adaptive explanations for such complex patterns invoke an interaction between selection for conspicuous signals and natural selection for crypsis. Although there is now a substantial body of evidence supporting the role of sexual selection for signaling functions, quantitative studies of crypsis remain comparatively rare. Here, we combine objective measures of coloration with information on predator visual sensitivities to study the role of crypsis in the evolution of color variation in an Australian lizard species complex (Ctenophorus decresii). We apply a model that allows us to quantify crypsis in terms of the visual contrast of the lizards against their natural backgrounds, as perceived by potential avian predators. We then use these quantitative estimates of crypsis to answer the following questions. Are there significant differences in crypsis conspicuousness among populations? Are there significant differences in crypsis conspicuousness between the sexes? Are body regions "exposed" to visual predators more cryptic than "hidden" body regions? Is there evidence for local adaptation with respect to crypsis against different substrates? In general, our results confirmed that there are real differences in crypsis conspicuousness both between populations and between sexes; that exposed body regions were significantly more cryptic than hidden ones, particularly in females; and that females, but not males, are more cryptic against their own local background than against the background of other populations. Body regions that varied most in contrast between the sexes and between populations were also most conspicuous and are emphasized by males during social and sexual signaling. However, results varied with respect to the aspect of coloration studied. Results based on chromatic contrast ("hue" of color) provided better support for the crypsis hypothesis than did results based on achromatic contrast ("brightness" of color). Taken together, these results support the view that crypsis plays a substantial role in the evolution of color variation and that color patterns represent a balance between the need for conspicuousness for signaling and the need for crypsis to avoid predation.
137 citations
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TL;DR: The sea level high, 120, 000 to 130, 000 years ago, did not lead to the complete inundation of Grand Cayman as discussed by the authors, however, it is evident that Cayman, Cayman Brac, and Little Cayman have not undergone independent vertical movement over the last 125 000 years.
Abstract: The sea level high, 120 000 to 130 000 years ago, did not lead to the complete inundation of Grand Cayman. Deposition associated with that highstand, which produced the limestones of the Ironshore Formation, occurred in small embayments on the south, east, and north coasts and in a large lagoon that covered the central and western parts of Grand Cayman. Limestones in the western part of that lagoon contain tidal (?) channels and large scour surfaces which resulted from storm activity. Overlying the lagoonal deposits are limestones that were deposited on a high-energy, prograding beach. These successions occur 2 to 7 m above present day sea level. Evidence obtained from wave-cut notches and the sedimentology of the Ironshore Formation on each of the Cayman Islands indicates that the late Pleistocene highstand was about 6 m above present day sea level. By comparing the present day elevations of the features used to determine this position, it is evident that Grand Cayman, Cayman Brac, and Little Cayman have not undergone independent vertical movement over the last 125 000 years.
54 citations
TL;DR: To investigate the significance of geographic variation in dewlap color, it was decided to describe it in greater detail across a series of Haitian populations of A. brevirostris and to determine the relationships of those same species.
Abstract: Seven to thirty ecologically diverse species of the iguanid genus Anolis live on each of the four Greater Antilles. Except quite locally, these species rarely occur without congeners. Usually an individual will encounter in a single day-or at least in its lifetime-individuals of from three to as many as six or seven other species. The dewlap, an extensible fold of skin contracted against the throat when not in use, is believed to be an important character by which these highly visual animals discriminate between members of their own and other species (Rand and Williams, 1970). Males of most anoles retain the ancestral condition of a large dewlap, though it is secondarily reduced or lost in a few species. By contrast, the dewlap of females is normally small. Dewlap extension is a component of displays in many anole species, but display repertoires and functions have been analyzed in only a few. There is evidence that displays involving the dewlap are used throughout the process of mate selection, which itself is poorly understood. A female may associate with a conspecific male on the basis of his common (assertion) display, with the male so selected enjoying the status of preferred mate when the female becomes receptive (Jenssen, 1970a, 1971). In several species, the dewlap is routinely extended in the male display that immediately precedes copulation (Greenberg and Noble, 1944; Gordon, 1956; Kastle, 1963; Rand, 1967; Ruibal, 1967); and while there are display patterns unique to this final stage of courtship, the common display may be given in combination with them (Kastle, 1963; Jenssen, 1970b). Indirect evidence further points to the importance of the dewlap as an isolating mechanism. Typically the dewlap of each member of a sympatric association is distinctive. For example, in each of the eight species known or suspected to occur at La Palma, Dominican Republic, the combination of colors differs; and size and shape also vary markedly (Rand and Williams, 1970). Pairs of closely related species often have widely divergent dewlap colors. Although geographic variation in dewlap color is not rare, intraspecific differences are generally less than those between sympatric species. However, two common Hispaniolan species, Anolis distichus and A. brevirostris, deviate in showing complex and striking geographic variation in dewlap color. Both species are relatively small and thick-bodied, differing in body color and pattern but not consistently in any scale character (Schwartz, 1968). In the lowlands of Hispaniola, A. distichus occupies mesic habitats, whereas A. brevirostris occurs in those that are more arid and therefore has a notably irregular range (Fig. 1). Even though these species do not overlap broadly, they do make contact: both have been found on the same tree in many areas. Major variation in dewlap color has been described and figured for A. distichus (Schwartz, 1968) and recorded for A. brevirostris (field notes of J. D. Lazell, A. S. Rand, A. Schwartz, and E. E. Williams). To investigate the significance of such variation, we decided to describe it in greater detail across a series of Haitian populations of A. brevirostris and to determine the relationships of those same
53 citations
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TL;DR: These findings suggest that acclimation of integumentary permeability may result from the gradual deposition of lipids in response to a local stimulus acting at the integument.
Abstract: Rates of cutaneous evaporative water loss (EWL) were measured in Anolis carolinensis lizards following their exposure to humid (80%-85% RH) or dry (35%-42% RH) environments. After 8 d of acclimation, lizards exposed to dry air had significantly reduced rates of EWL, whereas those exposed to humid air did not. Acclimation of EWL occurred regardless of whether lizards exposed to dry conditions were allowed to dehydrate (to 80% of initial mass) and regardless of whether lizards molted. Evaporation rates measured from isolated dorsal skin in vitro demonstrated that dry-acclimated lizards had a lower skin permeability than did humid-acclimated lizards. The differences in permeability disappeared when lipids were extracted from the skin. Our findings suggest that acclimation of integumentary permeability may result from the gradual deposition of lipids in response to a local stimulus acting at the integument.
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TL;DR: The first such evidence for lizards is reported, this in Anolis sagrei on Great Abaco Island, Bahamas, where the polymorphism occurs only in females, a restriction usual for insects but very rare in vertebrates.
Abstract: Polymorphism in body color and pattern occurs in an enormous diversity of animals, from crustaceans to mammals (Mayr, 1963). Its adaptive significance is often interpreted in terms of differential resemblance, protective or otherwise, to the animal's typical background. Yet substantial field evidence for this explanation exists only for a small fraction of species [for example, butterflies (Clarke and Sheppard, 1962; Curio, 1965); snails (Cain and Sheppard, 1954; Clarke, 1968; Owen, 1969; Arnold, 1969); frogs (Nevo, 1973); salamanders (Williams et al., 1968); snakes (Camin and Ehrlich, 1958); birds (Hall et al., 1966); and mammals (Gershenson, 1945) ]. Here we report the first such evidence for lizards, this in Anolis sagrei on Great Abaco Island, Bahamas. The polymorphism occurs only in females, a restriction usual for insects (Richards, 1961) but very rare in vertebrates; in fact, to our knowledge it occurs only in a handful of other Anolis and the partridge Lyrurus tetrix (Stegmann, 1932, cited by Huxley, 1955). In some vertebrates, however, polymorphism is more clearly expressed in females than males: e.g., the lizard Uta stansburiana (Ballinger and McKinney, 1967). We discuss possible reasons for this polymorphism's rarity after presenting the data.
48 citations
"Habitat light, colour variation, an..." refers background in this paper
...Second, females in many species of anoles exhibit a dorsal stripe or chevron pattern (e.g. Lazell, 1964; Schoener & Schoener, 1976) that almost certainly serves a disruptive coloration function (Cott, 1940; 0.0 700 1.0 Wavelength (nm) N or m al iz ed d ew la p re fl ec ta n…...
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